Classic papers and reading lists

The water maze sits on a long line of work about how animals, and people, find their way. Here are the papers worth knowing, grouped so you can start where your question is. Every paper listed has been checked against at least one bibliographic database, and we only show a summary where an editor has written and checked one.

The water maze itself

• Morris (1981). Spatial localization does not require the presence of local cues. Learning and Motivation 12(2):239-260. doi:10.1016/0023-9690(81)90020-5
  Morris showed that rats can learn to escape onto a platform hidden in opaque water using distal room cues, without any local cue marking the goal. This paper introduced the water maze as a test of spatial localisation.
• Morris et al. (1982). Place navigation impaired in rats with hippocampal lesions. Nature 297(5868):681-683. doi:10.1038/297681a0
  Morris, Garrud, Rawlins and O'Keefe found that rats with hippocampal lesions were impaired at finding a hidden platform in the water maze but could still swim to a visible platform. The study linked water maze place navigation to hippocampal function.
• Morris (1984). Developments of a water-maze procedure for studying spatial learning in the rat. Journal of Neuroscience Methods 11(1):47-60. doi:10.1016/0165-0270(84)90007-4
  Morris describes developments of the open-field water maze procedure in which rats learn to escape from opaque water onto a hidden platform. It covers automatic tracking of the rat's position without attached LEDs, and variants for studying different aspects of spatial memory (such as working memory) and non-spatial discrimination learning.
• Morris et al. (1986). Selective impairment of learning and blockade of long-term potentiation by an N-methyl-D-aspartate receptor antagonist, AP5. Nature 319(6056):774-776. doi:10.1038/319774a0
  Morris, Anderson, Lynch and Baudry showed that chronic infusion of the NMDA receptor antagonist AP5 impaired place learning in the water maze while blocking hippocampal long-term potentiation, without affecting visual discrimination learning. The study linked NMDA-dependent plasticity to spatial learning.
• Vorhees & Williams (2006). Morris water maze: procedures for assessing spatial and related forms of learning and memory. Nature Protocols 1(2):848-858. doi:10.1038/nprot.2006.116
  Vorhees and Williams give a detailed protocol for the Morris water maze, covering acquisition, probe (reference memory), reversal and cued trials. They explain how search patterns and cued trials help separate spatial learning from performance factors such as swimming ability or vision.
• Vorhees & Williams (2014). Assessing Spatial Learning and Memory in Rodents. ILAR Journal 55(2):310-332. doi:10.1093/ilar/ilu013
  Vorhees and Williams review methods for assessing spatial learning and memory in rodents, contrasting allocentric and egocentric navigation. The Morris water maze is treated in detail, alongside other mazes and their strengths and limitations.

The two original PDFs

The old watermaze.org hosted two papers as PDFs: Burešová and colleagues (1985) on the on-demand platform, and Markowska and colleagues (1993) on a variable-interval probe. Both PDFs are back at their original addresses (Buresova_JNM_1985.pdf and Markowska_BN_1993.pdf), and each paper has its own page with a plain-words summary and the publisher's DOI link.

Papers using HVS Image

Curious who else has tracked their animals with HVS Image? Rather than keep a list that goes stale, we point you to a live Google Scholar search for papers that mention HVS Image. It updates itself as new papers appear, so it's always worth a fresh look.

Technical reference documents

Looking for the methods behind the tracking and the measures? Our technical reference documents are a sample of papers describing methods used.

Reading lists by topic

• Cognitive maps & the hippocampusTolman, O'Keefe, place cells and grid cells.
• Ageing & diseaseThe water maze in ageing, dementia models, stress and injury.
• Measures & strategiesMeasuring and classifying what the animal did.
• Virtual & human navigationVirtual water mazes and how people find their way.
• Related testsBarnes, radial arm, elevated plus, open field and Atlantis.
• Burešová 1985The on-demand platform, forerunner of the Atlantis platform.
• Markowska 1993A variable-interval probe test of spatial memory.

For a longer list of studies that used one tracking system, see the publications citing HVS Image.